27 March, 2009

J.B.S. Haldane

John Burdon Sanderson Haldane was a British-born geneticist and evolutionary biologist. He was one of the founders (along with Ronald Fisher and Sewall Wright) of population genetics.

Haldane was born in Oxford to physiologist John Scott Haldane and Louisa Kathleen Haldane (née Trotter), and descended from an aristocratic intellectual Scottish family. His younger sister, Naomi, became a writer. His uncle was Richard Haldane, 1st Viscount Haldane, politician and one time Secretary of State for War; his aunt was the author Elizabeth Haldane. His father was a scientist, a philosopher and a Liberal, and his mother was a Conservative. Haldane took interest in his father’s work very early in his childhood.

He was educated at Eton and New College Oxford and served in the British Army during the First World War in the Black Watch regiment.

Between 1919 and 1922 he was a Fellow of New College, Oxford, then moved to Cambridge University, where he accepted a Readership in Biochemistry at Trinity College and taught there until 1932. During his nine years at Cambridge, Haldane worked on enzymes and genetics, particularly the mathematical side of genetics. Haldane wrote many popular essays on science that were eventually collected and published in 1927 in a volume entitled Possible Worlds.

He then accepted a position as Professor of Genetics and moved to University College London where he spent most of his academic career. Four years later he became the first Weldon Professor of Biometry at University College London.

In 1923, in a talk given in Cambridge, Haldane, foreseeing the exhaustion of coal for power generation in Britain, proposed a network of hydrogen-generating windmills. This is the first proposal of the hydrogen-based renewable energy economy.

In 1925, G. E. Briggs and Haldane derived a new interpretation of the enzyme kinetics law described by Victor Henri in 1903, different from the 1913 Michaelis-Menten equation. Leonor Michaelis and Maud Menten assumed that enzyme (catalyst) and substrate (reactant) are in fast equilibrium with their complex, which then dissociates to yield product and free enzyme. The Briggs-Haldane equation was of the same algebraic form, but their derivation is based on the quasi steady state approximation, that is the concentration(s) of intermediate complex(es) do(es) not change. As a result, the microscopic meaning of the "Michaelis Constant" (km) is different. Although commonly referring it as Michaelis-Menten kinetics, most of the current models actually use the Briggs-Haldane derivation.

Haldane made many contributions to human genetics and was one of the three major figures to develop the mathematical theory of population genetics. He is usually regarded as the third of these in importance, after R. A. Fisher and Sewall Wright. His greatest contribution was in a series of ten papers on "A Mathematical Theory of Natural and Artificial Selection" which was the major series of papers on the mathematical theory of natural selection. It treated many major cases for the first time, showing the direction and rates of changes of gene frequencies. It also pioneered in investigating the interaction of natural selection with mutation and with migration. Haldane's book, The Causes of Evolution (1932), summarized these results, especially in its extensive appendix. This body of work was a component of what came to be known as the "modern evolutionary synthesis", re-establishing natural selection as the premier mechanism of evolution by explaining it in terms of the mathematical consequences of Mendelian genetics.

Haldane introduced many quantitative approaches in biology such as in his essay On Being the Right Size. His contributions to theoretical population genetics and statistical human genetics included the first methods using maximum likelihood for estimation of human linkage maps, and pioneering methods for estimating human mutation rates. His was the first to calculate the mutational load caused by recurring mutations at a gene locus, and to introduce the idea of a "cost of natural selection".

Haldane is also known for an observation from his essay, On Being the Right Size, which Jane Jacobs and others have since referred to as Haldane's principle. This is that sheer size very often defines what bodily equipment an animal must have: "Insects, being so small, do not have oxygen-carrying bloodstreams. What little oxygen their cells require can be absorbed by simple diffusion of air through their bodies. But being larger means an animal must take on complicated oxygen pumping and distributing systems to reach all the cells." The conceptual metaphor to animal body complexity has been of use in energy economics and secession ideas.

Haldane was a keen experimenter, willing to expose himself to danger to obtain data. One experiment involving elevated levels of oxygen saturation triggered a fit which resulted in him suffering crushed vertebrae. In his decompression chamber experiments, he and his volunteers suffered perforated eardrums, but, as Haldane stated in What is Life, "the drum generally heals up; and if a hole remains in it, although one is somewhat deaf, one can blow tobacco smoke out of the ear in question, which is a social accomplishment."

In 1952, he received the Darwin Medal from the Royal Society. In 1956, he was awarded the Huxley Memorial Medal of the Royal Anthropological Institute. Among other awards, he received the Feltrinelli Prize, an Honorary Doctorate of Science, an Honorary Fellowship at New College, and the Kimber Award of the U.S. National Academy of Sciences. He was awarded the Linnean Society of London's prestigious Darwin-Wallace Medal in 1958.

Haldane became a socialist during World War I, supported the Spanish Republic during the Spanish Civil War and finally become a Communist. He was an enthusiastic, idealistic Marxist, and wrote many articles in the Communist Daily Worker. He was the chairman of the editorial board of the London edition for several years.

His vision of the Socialist principle can be considered pragmatic. In On being the right size, Haldane doubted that socialism could be operated on the scale of the British Empire or the United States or, implicitly, the Soviet Union: "while nationalization of certain industries is an obvious possibility in the largest of states, I find it no easier to picture a completely socialized British Empire or United States than an elephant turning somersaults or a hippopotamus jumping a hedge."

In 1937, Haldane became a Marxist and an open supporter of the Communist Party although not a member of the party. In 1938, he proclaimed enthusiastically that "I think that Marxism is true." He joined the Communist Party of Great Britain in 1942. The first edition of his children's book My Friend Mr. Leakey contained an avowal of his Party membership which was removed from later editions.

Events in the Soviet Union, such as the rise of anti-Mendelian agronomist Trofim Lysenko and the crimes of Stalin, may have caused him to break with the Party later in life, although he showed a partial support of Lysenko and Stalin. Pressed to speak out about the rise of Lysenkoism and the persecution of geneticists in the Soviet Union as anti-Darwinist and the denouncement of genetics as incompatible with dialectical materialism, Haldane shifted the focus to the United Kingdom and a criticism of the dependence of scientific research on financial patronage.

In 1941, Haldane wrote about the Soviet trial of his friend and fellow geneticist Nikolai Vavilov:

"The controversy among Soviet geneticists has been largely one between the academic scientist, represented by Vavilov and interested primarily in the collection of facts, and the man who wants results, represented by Lysenko. It has been conducted not with venom, but in a friendly spirit. Lysenko said (in the October discussions of 1939): 'The important thing is not to dispute; let us work in a friendly manner on a plan elaborated scientifically. Let us take up definite problems, receive assignments from the People's Commissariat of Agriculture of the USSR and fulfill them scientifically. Soviet genetics, as a whole, is a successful attempt at synthesis of these two contrasted points of view.'"

His ambiguous attitude toward the persecution of Vavilov was explainable by the atmosphere of the period, where the involvement in the Communist movement needed an all-or-nothing stand. His attitude changed dramatically at the end of World War II, when Lysenkoism reached a totalitarian influence in the Communist movement. He then become an explicit critic of the regime.

He left the Party in 1950, shortly after considering standing for Parliament as a Communist Party candidate. He continued to admire Stalin, describing him in 1962 as "a very great man who did a very good job."

The most famous of Haldane's many students, John Maynard Smith, shared his mixture of political and scientific interests to some extent, but broke away from the Communist Party in 1956.

Haldane's move to India, initially to the Indian Statistical Institute (ISI) was influenced by a number of factors. Officially he stated that his chief political reason was in response to the Suez Crisis. He wrote: "Finally, I am going to India because I consider that recent acts of the British Government have been violations of international law." His interest in India was also due to his interest in biological research, belief that the warm climate would do him good and that India offered him freedom and shared socialist dreams.

At the ISI, he headed the biometry unit and spent time researching a range of topics and guiding other researchers around him. He was keenly interested in inexpensive research and he wrote to Julian Huxley about his observations on Vanellus malabaricus boasting that he made them from the comfort of his backyard. Haldane took an interest in anthropology, human genetics and botany. He advocated the use of Vigna sinensis (cowpea) as a model for studying plant genetics. He took an interest in the pollination of the common weed Lantana camara. The quantitative study of biology was his main focus and he lamented that Indian universities forced those who took up biology to give up on an education in mathematics. Haldane took an interest in the study of floral symmetry. His wife, Helen Spurway, conducted studies on wild silk moths. He was also interested in Hinduism and after his arrival he became a vegetarian. Unable to get along with the director, P.C. Mahalanobis, Haldane resigned in February 1961 and moved to a newly established biometry unit in Orissa.

Haldane became an Indian citizen.

Haldane was a famous science popularizer. His essay, Daedalus; or, Science and the Future (1924), was remarkable in predicting many scientific advances but has been criticized for presenting a too idealistic view of scientific progress. Haldane’s book shows the effect of the separation between sexual life and pregnancy as a satisfactory one on human psychology and social life. The book was regarded as shocking science fiction at the time, being the first book about ectogenesis (the development of foetuses in artificial wombs) - "test tube babies", brought to life without sexual intercourse or pregnancy.

Haldane was a friend of the author Aldous Huxley, who parodied him in the novel "Antic Hay" (1923) as Shearwater, "the biologist too absorbed in his experiments to notice his friends bedding his wife". Haldane's discourse in Daedalus on ectogenesis was an influence on Huxley's Brave New World (1932) which features a eugenic society.

C. S. Lewis wrote much of his three interplanetary space novels, The Space Trilogy, in response to Haldane, whom Lewis considered to be an immoral man. Lewis modelled the character Weston, featured in the first two books, Out of the Silent Planet and Perelandra, on Haldane.

Haldane was one of those, along with Olaf Stapledon, Charles Kay Ogden, I. A. Richards, and H. G. Wells, whom Lewis accused of scientism, "the belief that the supreme moral end is the perpetuation of our own species, and that this is to be pursued even if, in the process of being fitted for survival, our species has to be stripped of all those things for which we value it—-of pity, of happiness, and of freedom." Shortly after the third book of the Ransom Trilogy appeared, J. B. S. Haldane criticised all three of them in an article entitled "Auld Hornie, F.R.S.". The title reflects the sarcastic tone of the article, Auld Hornie being the pet name given to the devil by the Scots and F.R.S. standing for "Fellow of the Royal Society". Lewis’s response, "A Reply to Professor Haldane", was never published during his lifetime and apparently never seen by Haldane. In it, Lewis claims that he was attacking scientism, not scientists, by challenging the view of some that the supreme goal of our species is to perpetuate itself at any expense.

Haldane died on 1 December 1964.